Foot and ankle functional morphology in anthropoid primates and Miocene hominoids A Dissertation Presented to the Faculty of the Graduate School at the University of Missouri In Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy By: Sharon Kuo Dr. Carol V. Ward, Dissertation Supervisor May 2020 Approval Page The undersigned, appointed by the dean of the Graduate School, have examined the dissertation entitled: Foot and ankle functional morphology in anthropoid primates and Miocene hominoids Presented by Sharon Kuo, a candidate for the degree of doctor of philosophy, and hereby certify that, in their opinion, it is worthy of acceptance. _______________________________________ Professor Carol Ward _______________________________________ Professor Kevin Middleton _______________________________________ Professor Casey Holliday _______________________________________ Professor Trent Guess _______________________________________ Professor Nicholas Gidmark Acknowledgments Acknowledgments First, I would like to thank Carol Ward for admitting me into this program and for your constant support over the past five years. Your enthusiasm for research and belief in your students are unparalleled and I am so lucky to have you in my corner. To the rest of my committee (Kevin Middleton, Casey Holliday, Trent Guess, Nick Gidmark): thank you for greatly improving this dissertation and for your patience throughout this process. In particular, thanks to Kevin for reminding me that it would all be ok and for always answering my stupid questions without making me feel stupid. Nick, I cannot thank you enough for always answering my XROMM questions over the years and for your constant support, beginning with the early phases of this project. This dissertation would not have been possible without the significant efforts of other people. To the Ross Lab at the University of Chicago, particularly Callum Ross, Michael Granatosky, and Myra Laird, thank you so much for working tirelessly on the XROMM data that made up a significant portion of this dissertation. To Biren Patel, Caley Orr, Sergio Almécija, Jerry DeSilva, and Ellie McNutt: thank you for your generosity with your scan data. I would like to give credit to Morgan Alwell for her help in cleaning the tarsal scans. I owe many thanks to the MU Life Sciences Fellowship for supporting me during my PhD and for Debbie Allen for being so helpful and for feeding all the Life Sciences Fellows. Thank you to Jerry DeSilva for getting me interested in research and for the continued lessons on feet and functional morphology over the years. The stellar IA faculty offered invaluable expertise and advice, as well as constant emotional support. Ian George, Henry Tsai, Elizabeth Moffett, Amanda Smolinsky, Sarah Peacock, and Kaleb Sellers: thank you for showing me what successful graduate students ii Acknowledgments look like when I first got here. To Emily Middleton and Faye McGechie, thank you for the insightful conversations, constant support, and truly heartwarming friendship. Sean Greer and Mel Boeyer, thanks for being familiar faces in this program and for being my oldest friends at Mizzou. I am so glad to have some of my Mercyhurst family here with me. Spiro Sullivan, Emily Lessner, and Austin Lawrence, thanks for your friendship and the numerous conversations that helped me in my research and in life. Ian Cost (and your better half), thanks for the nerdy science conversations, constant pep talks, confidence boosters when I didn’t really deserve them, and random bird identifications. Saturday morning study breakfasts at the Cost house helped make Columbia feel like home. Outside of IA, many people got me through my PhD. There are too many people to thank individually in this regard, so to all of you, thanks for being there for me to talk about things other than school, for reminding me that the world outside of this program indeed still exists, for being my travel buddies and pen pals, for editing (especially Luke A.), for telling me to calm down whenever I needed to, and for sending papers that I didn’t have access to but didn’t have time to ILL over the years. To Gracie Humaita and the Black Sheep, thank you for saving what is left of my sanity and also pulling me back into the world outside of this dissertation. Thank you to Ashley Hammond and KFFS for swooping in while I was in the darkest dissertation depths to remind me of why I love this field and the people with whom I am lucky enough to share it. To my family, thank you for forgiving missed events, trips, and phone calls so that I could finish this program and for your undying support. I couldn’t have done this without you. iii Table of Contents Table of Contents Acknowledgments............................................................................................................... ii List of Figures .................................................................................................................... vi List of Tables .....................................................................................................................xv Abstract ............................................................................................................................ xvi Chapter 1: Introduction ........................................................................................................1 Chapter 2: X-ray reconstruction of moving morphology (XROMM) of the talocrural, subtalar, calcaneocuboid, and talonavicular joints in Macaca mulatta .............................60 Introduction ........................................................................................................................60 Methods..............................................................................................................................69 Results ................................................................................................................................73 Discussion ..........................................................................................................................77 Conclusion .........................................................................................................................82 Chapter 3: Morphological variation of the calcaneus, talus, cuboid, and navicular in extant anthropoid primates ...............................................................................................109 Introduction ......................................................................................................................109 Methods............................................................................................................................119 Results ..............................................................................................................................123 Discussion ........................................................................................................................132 Conclusion .......................................................................................................................140 Chapter 4: Foot and ankle functional morphology of Miocene hominoids ....................184 Introduction ......................................................................................................................184 Methods............................................................................................................................189 Results ..............................................................................................................................193 Discussion ........................................................................................................................197 Conclusion .......................................................................................................................206 Chapter 5: Summary and Conclusions .............................................................................250 References ........................................................................................................................263 Appendix ..........................................................................................................................290 iv Table of Contents Vita ...................................................................................................................................296 v List of Figures List of Figures Figure 1-1 Articulated human foot ....................................................................................22 Figure 1-2 Joints and bones of interest for this study .......................................................23 Figure 1-3 Movements of the foot ....................................................................................24 Figure 1-4 Calcanei, tali, cuboids, and naviculars from Macaca, Pan, and Homo ..........25 Figure 1-5 Talocrural joint in Homo sapiens ....................................................................27 Figure 1-6 Morphological features of the primate talar trochlea ......................................28 Figure 1-7 Medial and lateral malleolar articular surfaces on the talus ............................28 Figure 1-8 Subtalar articulations on the calcaneus (left) and talus (right) ........................29 Figure 1-9 Talonavicular articulations on the talus and navicular ....................................29 Figure 1-10 Calcaneocuboid joint articulations on the calcaneus and cuboid ..................30 Figure